Three Keys to the Radiation of Angiosperms Into Freezing Environments

Total Page:16

File Type:pdf, Size:1020Kb

Three Keys to the Radiation of Angiosperms Into Freezing Environments LETTER doi:10.1038/nature12872 Three keys to the radiation of angiosperms into freezing environments Amy E. Zanne1,2, David C. Tank3,4, William K. Cornwell5,6, Jonathan M. Eastman3,4, Stephen A. Smith7, Richard G. FitzJohn8,9, Daniel J. McGlinn10, Brian C. O’Meara11, Angela T. Moles6, Peter B. Reich12,13, Dana L. Royer14, Douglas E. Soltis15,16,17, Peter F. Stevens18, Mark Westoby9, Ian J. Wright9, Lonnie Aarssen19, Robert I. Bertin20, Andre Calaminus15, Rafae¨l Govaerts21, Frank Hemmings6, Michelle R. Leishman9, Jacek Oleksyn12,22, Pamela S. Soltis16,17, Nathan G. Swenson23, Laura Warman6,24 & Jeremy M. Beaulieu25 Early flowering plants are thought to have been woody species to greater heights: as path lengths increase so too does resistance5. restricted to warm habitats1–3. This lineage has since radiated into Among extant strategies, the most efficient method of water delivery almost every climate, with manifold growth forms4. As angiosperms is through large-diameter water-conducting conduits (that is, vessels spread and climate changed, they evolved mechanisms to cope with and tracheids) within xylem5. episodic freezing. To explore the evolution of traits underpinning Early in angiosperm evolution they probably evolved larger conduits the ability to persist in freezing conditions, we assembled a large for water transport, especially compared with their gymnosperm cousins14. species-level database of growth habit (woody or herbaceous; 49,064 Although efficient in delivering water, these larger cells would have species), as well as leaf phenology (evergreen or deciduous), diameter impeded angiosperm colonization of regions characterized by episodic of hydraulic conduits (that is, xylem vessels and tracheids) and climate freezing14,15, as the propensity for freezing-induced embolisms (air bub- occupancies (exposure to freezing). To model the evolution of spe- bles produced during freeze/thaw events that block hydraulic pathways) cies’ traits and climate occupancies, we combined these data with an increases as conduit diameter increases5. Three evolutionary solutions unparalleled dated molecular phylogeny (32,223 species) for land liidae plants. Here we show that woody clades successfully movedintofreezing- Magno prone environments by either possessing transport networks of small 5 M o safe conduits and/or shutting down hydraulic function by dropping n o c leaves during freezing. Herbaceous species largely avoided freezing o ty le periods by senescing cheaply constructed aboveground tissue. Growth d e o 6 a n id e habit has long been considered labile , but we find that growth habit s a o e r r was less labile than climate occupancy. Additionally, freezing envir- e p u onments were largely filled by lineages that had already become herbs S or, when remaining woody, already had small conduits (that is, the trait evolvedbefore the climate occupancy). By contrast, most decidu- ous woody lineages had an evolutionary shift to seasonally shedding their leaves only after exposure to freezing (that is, the climate occu- pancy evolved before the trait). For angiosperms to inhabit novel cold environments they had to gain new structural and functional trait solutions; our results suggest that many of these solutions were probably acquired before their foray into the cold. Flowering plants (angiosperms) today grow in a vast range of envir- S u p e r onmental conditions, with this breadth probably related to their diverse a s t e 7 r i d morphology and physiology . However, early angiosperms are gen- a erally thought to have been woody and restricted to warm understory e habitats1–3. Debate continues about these assertions, in part because of Figure 1 | Time-calibrated maximum-likelihood estimate of the molecular the paucity of fossils and uncertainty in reconstructing habits for these 8–11 phylogeny for 31,749 species of seed plants. The four major angiosperm first representatives . Nevertheless, greater mechanical strength of lineages discussed in the text are highlighted: Monocotyledoneae (green), woody tissue would have made extended lifespans possible at a height Magnoliidae (blue), Superrosidae (brown) and Superasteridae (yellow). necessary to compete for light12,13. A major challenge resulting from Non-seed plant outgroups (that is, bryophytes, lycophytes and monilophytes) increased stature is that hydraulic systems must deliver water at tension were removed for the purposes of visualization. 1Department of Biological Sciences, George Washington University, Washington DC 20052, USA. 2Center for Conservation and Sustainable Development, Missouri Botanical Garden, St Louis, Missouri 63121, USA. 3Department of Biological Sciences, University of Idaho, Moscow, Idaho 83844, USA. 4Institute for Bioinformatics and Evolutionary Studies, University of Idaho, Moscow, Idaho 83844, USA. 5Department of Ecological Sciences, Systems Ecology, de Boelelaan 1085, 1081 HV Amsterdam, the Netherlands. 6Evolution & Ecology Research Centre, School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney, New South Wales 2052, Australia. 7Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, Michigan 48109, USA. 8Department of Zoology and Biodiversity Research Centre, University of British Columbia, Vancouver, British Columbia V6T1Z4, Canada. 9Department of Biological Sciences, Macquarie University, Sydney, New South Wales 2109, Australia. 10Department of Biology and the Ecology Center, Utah State University, Logan, Utah 84322, USA. 11Department of Ecology and Evolutionary Biology, University of Tennessee, Knoxville, Tennessee 37996, USA. 12Department of Forest Resources, University of Minnesota, St Paul, Minnesota 55108, USA. 13Hawkesbury Institute for the Environment, University of Western Sydney, Penrith, New South Wales 2751, Australia. 14Department of Earth and Environmental Sciences, Wesleyan University, Middletown, Connecticut 06459, USA. 15Department of Biology, University of Florida, Gainesville, Florida 32611, USA. 16Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, USA. 17Genetics Institute, University of Florida, Gainesville, Florida 32611, USA. 18Department of Biology, University of Missouri—St Louis, St Louis, Missouri 63121, USA. 19Department of Biology, Queen’s University, Kingston, Ontario K7L 3N6, Canada. 20Department of Biology, College of the Holy Cross, Worcester, Massachusetts 01610, USA. 21Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, United Kingdom. 22Polish Academy of Sciences, Institute of Dendrology, 62-035 Kornik, Poland. 23Department of Plant Biology and Ecology, Evolutionary Biology and Behavior, Program, Michigan State University, East Lansing, Michigan 48824, USA. 24Institute of Pacific Islands Forestry, USDA Forest Service, Hilo, Hawaii 96720, USA. 25National Institute for Mathematical & Biological Synthesis, University of Tennessee, Knoxville, Tennessee 37996, USA. 6 FEBRUARY 2014 | VOL 506 | NATURE | 89 ©2014 Macmillan Publishers Limited. All rights reserved RESEARCH LETTER seemingly arose to address the challenges of freezing: (1) woody species .0 uC across a species’ range; and ‘freezing exposed’, encountering withstood freezing temperatures without serious loss of hydraulic func- temperatures #0uC somewhere across a species’ range. This dichotomy tion by building safe water-transport networks consisting of small-diameter assumes that climate tracking through environmental changes is more conduits; (2) woody species shut down hydraulic function by becom- common than the evolution of climate occupancy; this is more likely to ing deciduous, dropping leaves during freezing periods; and (3) herb- be true if freezing exposure has a physiological cost in regions without aceous species largely avoided freezing by senescing cheaply constructed freezing21. Species were further distinguished by leaf phenology (deciduous aboveground tissue and overwintering, probably as seeds or underground or evergreen); conduit diameter (large $0.044 mm, or small ,0.044 mm; storage organs. However, the order in which angiosperms are likely to as 0.044 mm diameter is the diameter above which freezing-induced have acquired these solutions relative to exposure to and persistence in embolisms are believed to become frequent at modest tensions22); and the cold16 remains unclear. growth form (woody or herbaceous, with woody species defined as Proportions of herbaceous species, deciduous species and those with those maintaining a prominent aboveground stem that is persistent small water-conducting conduits increase towards the poles1,4,17,18, and over time and with changing environmental conditions; see Extended an earlier limited survey of angiosperm families indicated that herba- Data Fig. 1 for examples of angiosperms with woody growth habits as ceousness and ability to cope with freezing evolved in parallel19.However, we define them, and Extended Data Table 1 for a breakdown of growth exactly how global-scale ecological patterns are linked to functional evolu- habit by order within angiosperms). tion of angiosperms is uncertain. We dissect the contributions of different Among woody species we asked whether evolutionary transitions evolutionary solutions allowing angiosperms to cope with periodic freez- between climate occupancy states were significantly associated with shifts ing and assess likely pathways by which clades acquired these traits (that is, in leaf phenology and/or conduit diameter. Among all angiosperms we timing of evolution
Recommended publications
  • Toward a Resolution of Campanulid Phylogeny, with Special Reference to the Placement of Dipsacales
    TAXON 57 (1) • February 2008: 53–65 Winkworth & al. • Campanulid phylogeny MOLECULAR PHYLOGENETICS Toward a resolution of Campanulid phylogeny, with special reference to the placement of Dipsacales Richard C. Winkworth1,2, Johannes Lundberg3 & Michael J. Donoghue4 1 Departamento de Botânica, Instituto de Biociências, Universidade de São Paulo, Caixa Postal 11461–CEP 05422-970, São Paulo, SP, Brazil. [email protected] (author for correspondence) 2 Current address: School of Biology, Chemistry, and Environmental Sciences, University of the South Pacific, Private Bag, Laucala Campus, Suva, Fiji 3 Department of Phanerogamic Botany, The Swedish Museum of Natural History, Box 50007, 104 05 Stockholm, Sweden 4 Department of Ecology & Evolutionary Biology and Peabody Museum of Natural History, Yale University, P.O. Box 208106, New Haven, Connecticut 06520-8106, U.S.A. Broad-scale phylogenetic analyses of the angiosperms and of the Asteridae have failed to confidently resolve relationships among the major lineages of the campanulid Asteridae (i.e., the euasterid II of APG II, 2003). To address this problem we assembled presently available sequences for a core set of 50 taxa, representing the diver- sity of the four largest lineages (Apiales, Aquifoliales, Asterales, Dipsacales) as well as the smaller “unplaced” groups (e.g., Bruniaceae, Paracryphiaceae, Columelliaceae). We constructed four data matrices for phylogenetic analysis: a chloroplast coding matrix (atpB, matK, ndhF, rbcL), a chloroplast non-coding matrix (rps16 intron, trnT-F region, trnV-atpE IGS), a combined chloroplast dataset (all seven chloroplast regions), and a combined genome matrix (seven chloroplast regions plus 18S and 26S rDNA). Bayesian analyses of these datasets using mixed substitution models produced often well-resolved and supported trees.
    [Show full text]
  • Dipterocarps
    1682 TROPICAL ECOSYSTEMS / Dipterocarps Dipterocarps B Krishnapillay, Forest Research Institute Malaysia, Kepong, Malaysia & 2004, Elsevier Ltd. All Rights Reserved. Introduction The dipterocarp forests of Southeast Asia constitute a dominant and particularly valuable component of the world’s tropical rainforest. As a family of plants, Dipterocarpaceae may perhaps hold the distinction of being the best-known trees in the tropics. Their ecosystems are extremely diverse. They are uneven in their age and multilayered. They grow all the year round under warm temperatures and on sites where there is a large amount of rainfall. However, those growing in the seasonal forest are generally medium sized with the tallest trees being around 20 m with a maximum diameter of about 50 cm. Generally dipterocarps have been observed to occur on soils with very low fertility. Currently the dipterocarps dominate the international tropical timber market, and therefore play an important role in the economy of many Southeast Asian countries. In addition to Figure 1 Phytogeographical distribution of the family Diptero- timber, this family of trees also produces other non- carpaceae worldwide. timber products like resins and oleoresins. 3. South Asia, which constitutes India, the Andaman Distribution Islands, Bangladesh, and Nepal. The present distribution patterns of dipterocarps are 4. Sri Lanka. thought to reflect routes of colonization and past 5. The Seychelles. climatic conditions. They are distributed over the 6. Africa, which constitutes Madagascar, a narrow tropical belts of three continents of Asia, Africa, and strip from Mali to Sudan in the northern hemi- South America (Figure 1). They occupy several phyto- sphere, and Congo.
    [Show full text]
  • Well-Known Plants in Each Angiosperm Order
    Well-known plants in each angiosperm order This list is generally from least evolved (most ancient) to most evolved (most modern). (I’m not sure if this applies for Eudicots; I’m listing them in the same order as APG II.) The first few plants are mostly primitive pond and aquarium plants. Next is Illicium (anise tree) from Austrobaileyales, then the magnoliids (Canellales thru Piperales), then monocots (Acorales through Zingiberales), and finally eudicots (Buxales through Dipsacales). The plants before the eudicots in this list are considered basal angiosperms. This list focuses only on angiosperms and does not look at earlier plants such as mosses, ferns, and conifers. Basal angiosperms – mostly aquatic plants Unplaced in order, placed in Amborellaceae family • Amborella trichopoda – one of the most ancient flowering plants Unplaced in order, placed in Nymphaeaceae family • Water lily • Cabomba (fanwort) • Brasenia (watershield) Ceratophyllales • Hornwort Austrobaileyales • Illicium (anise tree, star anise) Basal angiosperms - magnoliids Canellales • Drimys (winter's bark) • Tasmanian pepper Laurales • Bay laurel • Cinnamon • Avocado • Sassafras • Camphor tree • Calycanthus (sweetshrub, spicebush) • Lindera (spicebush, Benjamin bush) Magnoliales • Custard-apple • Pawpaw • guanábana (soursop) • Sugar-apple or sweetsop • Cherimoya • Magnolia • Tuliptree • Michelia • Nutmeg • Clove Piperales • Black pepper • Kava • Lizard’s tail • Aristolochia (birthwort, pipevine, Dutchman's pipe) • Asarum (wild ginger) Basal angiosperms - monocots Acorales
    [Show full text]
  • Human-Mediated Introductions of Australian Acacias
    Diversity and Distributions, (Diversity Distrib.) (2011) 17, 771–787 S EDITORIAL Human-mediated introductions of PECIAL ISSUE Australian acacias – a global experiment in biogeography 1 2 1 3,4 David M. Richardson *, Jane Carruthers , Cang Hui , Fiona A. C. Impson , :H Joseph T. Miller5, Mark P. Robertson1,6, Mathieu Rouget7, Johannes J. Le Roux1 and John R. U. Wilson1,8 UMAN 1 Centre for Invasion Biology, Department of ABSTRACT - Botany and Zoology, Stellenbosch University, MEDIATED INTRODUCTIONS OF Aim Australian acacias (1012 recognized species native to Australia, which were Matieland 7602, South Africa, 2Department of History, University of South Africa, PO Box previously grouped in Acacia subgenus Phyllodineae) have been moved extensively 392, Unisa 0003, South Africa, 3Department around the world by humans over the past 250 years. This has created the of Zoology, University of Cape Town, opportunity to explore how evolutionary, ecological, historical and sociological Rondebosch 7701, South Africa, 4Plant factors interact to affect the distribution, usage, invasiveness and perceptions of a Protection Research Institute, Private Bag globally important group of plants. This editorial provides the background for the X5017, Stellenbosch 7599, South Africa, 20 papers in this special issue of Diversity and Distributions that focusses on the 5Centre for Australian National Biodiversity global cross-disciplinary experiment of introduced Australian acacias. A Journal of Conservation Biogeography Research, CSIRO Plant Industry, GPO Box Location Australia and global. 1600, Canberra, ACT, Australia, 6Department of Zoology and Entomology, University of Methods The papers of the special issue are discussed in the context of a unified Pretoria, Pretoria 0002, South Africa, framework for biological invasions.
    [Show full text]
  • Plastid Phylogenomic Insights Into the Evolution of the Caprifoliaceae S.L. (Dipsacales)
    Molecular Phylogenetics and Evolution 142 (2020) 106641 Contents lists available at ScienceDirect Molecular Phylogenetics and Evolution journal homepage: www.elsevier.com/locate/ympev Plastid phylogenomic insights into the evolution of the Caprifoliaceae s.l. T (Dipsacales) Hong-Xin Wanga,1, Huan Liub,c,1, Michael J. Moored, Sven Landreine, Bing Liuf,g, Zhi-Xin Zhua, ⁎ Hua-Feng Wanga, a Key Laboratory of Tropical Biological Resources of Ministry of Education, School of Life and Pharmaceutical Sciences, Hainan University, Haikou 570228, China b BGI-Shenzhen, Beishan Industrial Zone, Yantian District, Shenzhen 518083, China c State Key Laboratory of Agricultural Genomics, BGI-Shenzhen, Shenzhen 518083, China d Department of Biology, Oberlin College, Oberlin, OH 44074, USA e Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Menglun, 666303, China f State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Science, Beijing 100093, China g Sino-African Joint Research Centre, Chinese Academy of Science, Wuhan 430074, China ARTICLE INFO ABSTRACT Keywords: The family Caprifoliaceae s.l. is an asterid angiosperm clade of ca. 960 species, most of which are distributed in Caprifoliaceae s.l. temperate regions of the northern hemisphere. Recent studies show that the family comprises seven major Dipsacales clades: Linnaeoideae, Zabelia, Morinoideae, Dipsacoideae, Valerianoideae, Caprifolioideae, and Diervilloideae. Plastome However, its phylogeny at the subfamily or genus level remains controversial, and the backbone relationships Phylogenetics among subfamilies are incompletely resolved. In this study, we utilized complete plastome sequencing to resolve the relationships among the subfamilies of the Caprifoliaceae s.l. and clarify several long-standing controversies. We generated and analyzed plastomes of 48 accessions of Caprifoliaceae s.l., representing 44 species, six sub- families and one genus.
    [Show full text]
  • The Origin of the Bifurcating Style in Asteraceae (Compositae)
    Annals of Botany 117: 1009–1021, 2016 doi:10.1093/aob/mcw033, available online at www.aob.oxfordjournals.org The origin of the bifurcating style in Asteraceae (Compositae) Liliana Katinas1,2,*, Marcelo P. Hernandez 2, Ana M. Arambarri2 and Vicki A. Funk3 1Division Plantas Vasculares, Museo de La Plata, La Plata, Argentina, 2Laboratorio de Morfologıa Comparada de Espermatofitas (LAMCE), Facultad de Ciencias Agrarias y Forestales, Universidad Nacional de La Plata, La Plata, Argentina and 3Department of Botany, NMNH, Smithsonian Institution, Washington D.C., USA *For correspondence. E-mail [email protected] Received: 20 November 2015 Returned for revision: 22 December 2015 Accepted: 8 January 2016 Published electronically: 20 April 2016 Background and Aims The plant family Asteraceae (Compositae) exhibits remarkable morphological variation in the styles of its members. Lack of studies on the styles of the sister families to Asteraceae, Goodeniaceae and Calyceraceae, obscures our understanding of the origin and evolution of this reproductive feature in these groups. The aim of this work was to perform a comparative study of style morphology and to discuss the relevance of im- portant features in the evolution of Asteraceae and its sister families. Methods The histochemistry, venation and general morphology of the styles of members of Goodeniaceae, Calyceraceae and early branching lineages of Asteraceae were analysed and put in a phylogenetic framework to dis- cuss the relevance of style features in the evolution of these families. Key Results The location of lipophilic substances allowed differentiation of receptive from non-receptive style papillae, and the style venation in Goodeniaceae and Calyceraceae proved to be distinctive.
    [Show full text]
  • Phylogeny of the Tropical Tree Family Dipterocarpaceae Based on Nucleotide Sequences of the Chloroplast Rbcl Gene1
    American Journal of Botany 86(8): 1182±1190. 1999. PHYLOGENY OF THE TROPICAL TREE FAMILY DIPTEROCARPACEAE BASED ON NUCLEOTIDE SEQUENCES OF THE CHLOROPLAST RBCL GENE1 S. DAYANANDAN,2,6 PETER S. ASHTON,3 SCOTT M. WILLIAMS,4 AND RICHARD B. PRIMACK2 2Biology Department, Boston University, Boston, Massachusetts 02215; 3Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138; and 4Division of Biomedical Sciences, Meharry Medical College, 1005 D. B. Todd, Jr. Boulevard, Nashville, Tennessee 37208 The Dipterocarpaceae, well-known trees of the Asian rain forests, have been variously assigned to Malvales and Theales. The family, if the Monotoideae of Africa (30 species) and South America and the Pakaraimoideae of South America (one species) are included, comprises over 500 species. Despite the high diversity and ecological dominance of the Dipterocar- paceae, phylogenetic relationships within the family as well as between dipterocarps and other angiosperm families remain poorly de®ned. We conducted parsimony analyses on rbcL sequences from 35 species to reconstruct the phylogeny of the Dipterocarpaceae. The consensus tree resulting from these analyses shows that the members of Dipterocarpaceae, including Monotes and Pakaraimaea, form a monophyletic group closely related to the family Sarcolaenaceae and are allied to Malvales. The present generic and higher taxon circumscriptions of Dipterocarpaceae are mostly in agreement with this molecular phylogeny with the exception of the genus Hopea, which forms a clade with Shorea sections Anthoshorea and Doona. Phylogenetic placement of Dipterocarpus and Dryobalanops remains unresolved. Further studies involving repre- sentative taxa from Cistaceae, Elaeocarpaceae, Hopea, Shorea, Dipterocarpus, and Dryobalanops will be necessary for a comprehensive understanding of the phylogeny and generic limits of the Dipterocarpaceae.
    [Show full text]
  • High Tree Endemism Recorded in Kanana Kanda Isolated Forest Fragment in Wet Zone of Sri Lanka
    International Journal of Agriculture, Forestry and Plantation, Vol. 2 (February.) ISSN 2462-1757 2 01 6 HIGH TREE ENDEMISM RECORDED IN KANANA KANDA ISOLATED FOREST FRAGMENT IN WET ZONE OF SRI LANKA P.K.J. De Mel Department of Agricultural and Plantation Engineering Open University of Sri Lanka, P.O. Box 21, Nawala, Nugegoda (10250), Sri Lanka Email: [email protected] K.A.J.M. Kuruppuarachchi Department of Botany Open University of Sri Lanka, P.O. Box 21, Nawala, Nugegoda (10250), Sri Lanka Email: [email protected] ABSTRACT Kanana Kanda is an isolated lowland hill with an altitude of 115m covered by natural forest with an extent of 13ha. The forest fragment located in wet zone of Sri Lanka where much species diversity and endemism is found. The forest is disappearing fast due to anthropogenic influences. Therefore the present study was carried out with the objective of assessment of existing tree flora. Reconnaissance survey was first conducted in the forest in order to gather basic information on vegetation types and floristic characteristics. Four transects with a size of 100m x 5m each were laid for sampling trees. A woody plant with a dbh equal or greater than 5cm considered as a tree. A total number of 464 trees were enumerated in the forest. Field identification of species performed with the consultation of personnel who have sufficient knowledge, skills and experience on similar vegetation. Herbarium specimens were prepared from each tree unidentified in the field and later identified those comparing with the specimens preserved in the National Herbarium. Present study, recorded a total number of 50 different species belongs to 29 families.
    [Show full text]
  • GENOME EVOLUTION in MONOCOTS a Dissertation
    GENOME EVOLUTION IN MONOCOTS A Dissertation Presented to The Faculty of the Graduate School At the University of Missouri In Partial Fulfillment Of the Requirements for the Degree Doctor of Philosophy By Kate L. Hertweck Dr. J. Chris Pires, Dissertation Advisor JULY 2011 The undersigned, appointed by the dean of the Graduate School, have examined the dissertation entitled GENOME EVOLUTION IN MONOCOTS Presented by Kate L. Hertweck A candidate for the degree of Doctor of Philosophy And hereby certify that, in their opinion, it is worthy of acceptance. Dr. J. Chris Pires Dr. Lori Eggert Dr. Candace Galen Dr. Rose‐Marie Muzika ACKNOWLEDGEMENTS I am indebted to many people for their assistance during the course of my graduate education. I would not have derived such a keen understanding of the learning process without the tutelage of Dr. Sandi Abell. Members of the Pires lab provided prolific support in improving lab techniques, computational analysis, greenhouse maintenance, and writing support. Team Monocot, including Dr. Mike Kinney, Dr. Roxi Steele, and Erica Wheeler were particularly helpful, but other lab members working on Brassicaceae (Dr. Zhiyong Xiong, Dr. Maqsood Rehman, Pat Edger, Tatiana Arias, Dustin Mayfield) all provided vital support as well. I am also grateful for the support of a high school student, Cady Anderson, and an undergraduate, Tori Docktor, for their assistance in laboratory procedures. Many people, scientist and otherwise, helped with field collections: Dr. Travis Columbus, Hester Bell, Doug and Judy McGoon, Julie Ketner, Katy Klymus, and William Alexander. Many thanks to Barb Sonderman for taking care of my greenhouse collection of many odd plants brought back from the field.
    [Show full text]
  • Dry Forest Trees of Madagascar
    The Red List of Dry Forest Trees of Madagascar Emily Beech, Malin Rivers, Sylvie Andriambololonera, Faranirina Lantoarisoa, Helene Ralimanana, Solofo Rakotoarisoa, Aro Vonjy Ramarosandratana, Megan Barstow, Katharine Davies, Ryan Hills, Kate Marfleet & Vololoniaina Jeannoda Published by Botanic Gardens Conservation International Descanso House, 199 Kew Road, Richmond, Surrey, TW9 3BW, UK. © 2020 Botanic Gardens Conservation International ISBN-10: 978-1-905164-75-2 ISBN-13: 978-1-905164-75-2 Reproduction of any part of the publication for educational, conservation and other non-profit purposes is authorized without prior permission from the copyright holder, provided that the source is fully acknowledged. Reproduction for resale or other commercial purposes is prohibited without prior written permission from the copyright holder. Recommended citation: Beech, E., Rivers, M., Andriambololonera, S., Lantoarisoa, F., Ralimanana, H., Rakotoarisoa, S., Ramarosandratana, A.V., Barstow, M., Davies, K., Hills, BOTANIC GARDENS CONSERVATION INTERNATIONAL (BGCI) R., Marfleet, K. and Jeannoda, V. (2020). Red List of is the world’s largest plant conservation network, comprising more than Dry Forest Trees of Madagascar. BGCI. Richmond, UK. 500 botanic gardens in over 100 countries, and provides the secretariat to AUTHORS the IUCN/SSC Global Tree Specialist Group. BGCI was established in 1987 Sylvie Andriambololonera and and is a registered charity with offices in the UK, US, China and Kenya. Faranirina Lantoarisoa: Missouri Botanical Garden Madagascar Program Helene Ralimanana and Solofo Rakotoarisoa: Kew Madagascar Conservation Centre Aro Vonjy Ramarosandratana: University of Antananarivo (Plant Biology and Ecology Department) THE IUCN/SSC GLOBAL TREE SPECIALIST GROUP (GTSG) forms part of the Species Survival Commission’s network of over 7,000 Emily Beech, Megan Barstow, Katharine Davies, Ryan Hills, Kate Marfleet and Malin Rivers: BGCI volunteers working to stop the loss of plants, animals and their habitats.
    [Show full text]
  • Reconstructing the Basal Angiosperm Phylogeny: Evaluating Information Content of Mitochondrial Genes
    55 (4) • November 2006: 837–856 Qiu & al. • Basal angiosperm phylogeny Reconstructing the basal angiosperm phylogeny: evaluating information content of mitochondrial genes Yin-Long Qiu1, Libo Li, Tory A. Hendry, Ruiqi Li, David W. Taylor, Michael J. Issa, Alexander J. Ronen, Mona L. Vekaria & Adam M. White 1Department of Ecology & Evolutionary Biology, The University Herbarium, University of Michigan, Ann Arbor, Michigan 48109-1048, U.S.A. [email protected] (author for correspondence). Three mitochondrial (atp1, matR, nad5), four chloroplast (atpB, matK, rbcL, rpoC2), and one nuclear (18S) genes from 162 seed plants, representing all major lineages of gymnosperms and angiosperms, were analyzed together in a supermatrix or in various partitions using likelihood and parsimony methods. The results show that Amborella + Nymphaeales together constitute the first diverging lineage of angiosperms, and that the topology of Amborella alone being sister to all other angiosperms likely represents a local long branch attrac- tion artifact. The monophyly of magnoliids, as well as sister relationships between Magnoliales and Laurales, and between Canellales and Piperales, are all strongly supported. The sister relationship to eudicots of Ceratophyllum is not strongly supported by this study; instead a placement of the genus with Chloranthaceae receives moderate support in the mitochondrial gene analyses. Relationships among magnoliids, monocots, and eudicots remain unresolved. Direct comparisons of analytic results from several data partitions with or without RNA editing sites show that in multigene analyses, RNA editing has no effect on well supported rela- tionships, but minor effect on weakly supported ones. Finally, comparisons of results from separate analyses of mitochondrial and chloroplast genes demonstrate that mitochondrial genes, with overall slower rates of sub- stitution than chloroplast genes, are informative phylogenetic markers, and are particularly suitable for resolv- ing deep relationships.
    [Show full text]
  • Anatomy and Affinities of Penthorum Melanie Lynn Haskins
    University of Richmond UR Scholarship Repository Biology Faculty Publications Biology 2-1987 Anatomy and Affinities of Penthorum Melanie Lynn Haskins W. John Hayden University of Richmond, [email protected] Follow this and additional works at: http://scholarship.richmond.edu/biology-faculty-publications Part of the Botany Commons, Other Plant Sciences Commons, and the Plant Biology Commons Recommended Citation Haskins, Melanie Lynn, and W. John Hayden. "Anatomy and Affinities of Penthorum." American Journal of Botany 74, no. 2 (February 1987): 164-77. This Article is brought to you for free and open access by the Biology at UR Scholarship Repository. It has been accepted for inclusion in Biology Faculty Publications by an authorized administrator of UR Scholarship Repository. For more information, please contact [email protected]. Amer. J. Bot. 74(2): 164-177. 1987. ANATOMY AND AFFINITIES OF PENTHORUM' MELANIE L. HASKINS AND W. JOHN HAYDEN Department of Biology, University of Richmond, Richmond, Virginia 23173 ABSTRACT The genus Penthorum L. consists of two species of perennial herbs, P. sedoides of eastern North America and P. chinense ofeastern Asia. Pentho rum has long been considered intermediate between Crassulaceae and Saxifragaceae. An anatomical study of both species was undertaken to contribute to a better understanding of the relationships ofthese plants. Prominent anatomical features of Penthorum include: an aerenchymatous cortex and closely-spaced collateral vascular bundles of stems; one-trace unilacunar nodes; brochidodromous venation, rosoid teeth bearing hydathodes, and anomocytic stomata of leaves; angular vessel elements with many-barred scalariform perforation plates and alternate to scattered intervascular pits; thin-walled non­ septate fiber-tracheids; abundant homocellular erect uniseriate and biseriate rays; and absence of axial xylem parenchyma.
    [Show full text]